The Grouse in Health and in Disease/Introduction

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By Lord Lovat

Before the formal appointment of the Committee in 1905 the following preliminary work of organisation was carried out.

On June 5th, 1904, the organisers of the present investigation met, and after discussion formed a Committee of Inquiry to investigate the subject of "Grouse Disease." The following gentlemen were present: The Marquis of Tullibardine, Lord Lovat, Mackintosh of Mackintosh, Mr R. H. Rimington Wilson, Mr J. Graham, Mr D. W. Drummond, Mr R. C. Munro Ferguson.

Lord Lovat was appointed Chairman, and Lord Onslow, the then President of the Board of Agriculture, was approached with the view of obtaining the assistance of that Board.

A further meeting was held in December of the same year, when the details of the proposed lines of inquiry were discussed, a Secretary was appointed, and a number of witnesses were examined. The formal appointment of the Committee as a Departmental Committee of the Board of Agriculture and Fisheries was intimated by the Secretary of the Department on April 13th, 1905. The terms of the appointment marked a departure from the usual procedure in such matters, for they provided that no public funds should be devoted to the Inquiry, but that the investigation should be conducted at the expense either of the members of the Committee or of private subscribers. The members included the above-named gentlemen, with the addition of Earl de Grey (now Marquis of Ripon) and Lord Henry Scott. Dr William Somerville was appointed to represent the Board of Agriculture and Fisheries, and upon his retirement from the Board Mr T. H. Middleton was appointed. The Committee sustained a severe loss by the death in 1910 of Mr James Graham, one of its most active and capable members.

In April and May 1905 an appeal was sent to a limited number of proprietors and tenants of Grouse moors asking for financial support. This appeal resulted in subscriptions amounting to over £400; these subscriptions were limited to a sum not exceeding £5 a year, and in the majority of cases were guaranteed for a period of three years. On the strength of this response a number of scientific gentlemen were asked to assist in the investigation, and a body of local correspondents in different parts of the country was appointed to make observations and to report upon any special local conditions or circumstances affecting Grouse in their respective districts. These local correspondents consisted mainly of resident proprietors, factors, estate agents, and gamekeepers. Great care was taken in their selection, and experience has shown that they have fully justified their appointment. About three hundred correspondents were formally appointed, and many other proprietors and gamekeepers corresponded regularly with the Secretary and with the staff of the Committee whenever occasion arose. The list of local correspondents might easily have been doubled by adding to it the names of those who had shown themselves able and willing to assist the investigation, but unfortunately the funds of the Committee would not admit of such addition. Lists of the Committee, of the staff, and of the local correspondents are given in Appendix A.

For the instruction of local correspondents and others who wished to be informed of the existing state of knowledge on the subject of "Grouse Disease," and further to indicate the exact points upon which information was required, the Committee drew up an illustrated pamphlet entitled "Notes on the Grouse"; in this a short summary was given of the life history of the bird, with a description of the typical characteristics of "Grouse Disease" as then recognised. The pamphlet called attention to the many theories which existed on the subject, and indicated the lines upon which the Committee proposed to carry out their investigation. This pamphlet was privately circulated among correspondents and subscribers, but was not offered for sale.

The scientific experts drew attention to the difficulty of carrying out experiments upon Grouse in a wild state, and accordingly in 1906 the Committee established an observation area in Surrey, where it was soon demonstrated that Grouse could be kept in captivity. The necessary licence was obtained from the Home Office. This observation area has been of the utmost value to the Committee.

Owing to the necessity of having a constant supply of healthy Grouse for examination in every month of the year, to enable the Committee to collect accurate information on the question of feeding, moulting, and seasonal changes, arrangements were made by members of the Committee and certain local correspondents to send to the Field Observer each monthly of the year a certain number of freshly killed birds. Many hundreds of such birds have been examined, and from the material so obtained valuable, and in many cases new information was gained. An interesting collection of over six hundred Grouse skins has been prepared, showing the types of plumage found in both sexes at different times of the year and in different districts, and also certain abnormalities. Selections from this collection and from the other material collected by the Committee were exhibited at a soirée of the Royal Society in May 1909, and at the Vienna Sports Exhibition of 1910.

The Committee began their observations in the field in the autumn of 1905; during this season and 1906 the stock of Grouse both in Scotland and England was remarkably healthy, and an excellent opportunity was thus given to study the bird under normal conditions. The Field Observer visited many moors, his visits extending over a period of seven months, from April to October. During this time he got into close touch with the Committee's correspondents in different parts of the country, checked their information, and with their assistance studied the varying conditions governing particular districts. Whenever a case of suspected "Grouse Disease" was reported the moor was visited by the Field Observer or one of his assistants, and specimens of suspicious birds were subjected to laboratory examination.

During 1907 a considerable mortality amongst the Grouse in certain districts was reported in the spring and early summer months. The Committee's experts made a very careful investigation into every case reported, but, contrary to expectation, it was not found that the character of the disease differed materially in its essential features from those occasional isolated cases of mortality which had occurred in the previous year. The Committee found no examples of the acute or sudden form of disease which had been described by former observers. The outbreak of mortality, however, gave an excellent opportunity for collecting data regarding the lingering or pining form of disease which has since been traced to the ravages of the threadworm Trichostrongylus pergracilis.[1]

By 1908 the Committee had completed the preliminary work required to enable the subject to be developed on scientific lines. Evidence and statistics had been collected which indicated the special directions in which further investigations were necessary or likely to be helpful. The natural history of the normal healthy Grouse had been fully studied, and the general pathological characteristics of "Grouse Disease," from a field observer's point of view, had been ascertained. Even at this date the Committee were of opinion that they had discovered the principal causes of mortality amongst Grouse; but until they had further confirmed their suspicions they decided not to publish anything in the nature of results. It was at this stage that an impatient public and the necessity to stimulate dilatory subscribers forced upon the Committee the necessity of publishing some account of their progress, and the Interim Report issued in August 1908 was the outcome of this demand. The Interim Report contained an account of the work done by the Committee up to date, but omitted all reference to the results which had only been achieved in part.

During the second or research stage of the investigation the following special points were studied: (l) The life history of the Trichostrongylus pergracilis, which the Committee believed to be the immediate cause of "Grouse Disease"; (2) The life history of the other internal parasites of Grouse; (3) The protozoal parasites infecting the alimentary tract and blood of Grouse; (4) The bacteriology of Grouse; (5) The various insects found on the moors both from the point of view of insect-borne disease and from the point of view of food; (6) The questions afiecting the food supply of Grouse, including the management of heather land, causes of destruction of heather, e.g., frost, heather-beetle, etc.

These lines of research were diligently followed up by the members of the Committee's Scientific Staff during the last three years of the Inquiry—the work entailed long series of experiments carried out upon the open moor, in the laboratory, or at the Frimley observation area. The results have been unexpectedly conclusive, considering the short time available for so great a task.

The Committee consider that although their immediate object has been achieved, viz., the elucidation of the causes of "Grouse Disease," the present Inquiry has scarcely crossed the threshold of the investigation into the general pathology of birds, and there is still a large amount of work which might be profitably under- taken. The most important department of the research, so far at least as relates to mortality amongst adult Grouse, was the investigation of the life history of the strongyle threadworm. The work was rendered difficult owing to the small size of this parasite, but thanks to the efforts of Dr Wilson, Dr Shipley, and Dr Leiper, we are now in a position to speak with something approaching certainty on the subject. These gentlemen have worked at the subject for more than three years, and have not only ascertained the life cycle through which this worm passes, but have discovered the conditions which are favourable or prejudicial to its growth; they have been able to rear the young strongyle, and by administering it through the medium of food to hand-reared Grouse free from nematode infection, have infected the hand-reared birds with " Grouse Disease."

Another interesting and important outcome of the Inquiry has been the discovery of a cause of death among Grouse in their infancy due to the presence of an intestinal parasite known as Eimeria (Coccidium) avium. It is unfortunate that the Inquiry is being brought to a close while Dr Fantham is still engaged in tracing the predisposing causes of this disease with a view to finding whether any preventive measures are possible. It is true that such preventive measures, even when found, might not be easily applied to the Grouse in a wild state; but they would be of the greatest possible value for the treatment of hand-reared game-birds or domestic fowls.

During the progress of the Inquiry many questions affecting Grouse and Grouse shooting, but not directly connected with disease, have come before the notice of the Committee, and owing to their general interest to readers of the Report it has been thought well to refer to some of them. Since the Inquiry has been mainly supported by those whose interests are more intimately connected with sport than with science, the inclusion of chapters on such subjects of practical importance as Moor Management, Heather-burning, Vermin, Keepers and the Value of Grouse moors, requires no apology. The chapters of natural history, such as Life History, Plumage changes, Food, Physiology and Death from Causes other than Disease, are all indirectly connected with the main subjects of the Inquiry.

It will be seen that by the inclusion of the above-mentioned chapters the Report of the Committee becomes a monograph on the Red Grouse in health and disease rather than a summary of the proceedings of a Departmental Committee of Inquiry.

During the period of the Inquiry a large number of Pamphlets, Reports, and Letters of Instructions have been printed and issued by the Committee to its local correspondents and other supporters. These documents, in addition to the "Notes on the Grouse" pamphlet already referred to, include Notes of Evidence taken at the meetings of Committee, Lists of Queries, Forms of Particulars of Specimens, Periodical Reports on the Progress of the Inquiry, Lists of Subscribers, Lists of Local Correspondents, Statements of Crop-contents, Circular Letters to Proprietors, etc., etc. In all more than 40,000 printed documents have been circulated, in addition to a large number of typewritten circulars and letters, of which no accurate record has been kept.

The correspondence both of the Secretary and the Field Observer has been voluminous, and has sometimes been subject to such sudden bursts of activity that it was found well-nigh impossible to keep pace with it. To this cause must be ascribed occasional failures to acknowledge written communications by return of post, for which failures the Committee now tender their apology.

In the course of the investigation many technical questions arose which made it necessary to employ the services of leading scientific experts, and, owing to the difficulty in obtaining immediate and definite results, it was found that the period of the Inquiry would have to be extended beyond the three years originally fixed. The result has been that the Committee found it necessary to exceed their original estimates.

During the whole Inquiry the Committee has been greatly hampered in their labours by lack of funds. The total income has never amounted to £1,000 in any one year, and the work would have been in danger of coming to an end were it not that many members of the Scientific Staff have given their services gratuitously or for at most a nominal consideration.

What success the Committee have met with is due to several causes. Firstly, the work was, in the main, directed by small Sub-Committees who were unhampered by official restrictions and untrammelled by traditional red tape. Secondly, the Chairman and the Secretary had the cordial support not only of the other members of the Committee but of all those directly or indirectly interested in the Grouse. Thirdly, the members of the Scientific Staff" took the keenest interest in the problems they sought to solve, and were willing to place their knowledge, their spare time, and their technical skill at the disposal of the Committee unremunerated, or at best remunerated at an entirely inadequate scale. Fourthly, the Inquiry aroused a certain public spirit, which not only found expression in the willingness of sportsmen, landlords, keepers, and others to do all in their power to assist the work of the Committee, but led the printers, the firm which supplied the paper upon which the book is printed, the publishers and many others connected with the preparation of the volume, to grant the Committee the most favourable terms.

That this Inquiry did not cost more than the small sum of £4,366 in the six years over which the work extended (averaging £727 a year) is due to the causes set forth above, and to the constant vigilance and unselfish insistance on economy on the part of the Secretary. Compared with the cost of similar Royal Commissions and Departmental Committees this sum is a mere trifle, but it shows that satisfactory results can be attained at very small expense. Much money was of course saved by not printing the evidence given at the numerous examinations of gamekeepers and others held by the Committee. Such evidence is, as a rule, printed in full, and remains unheeded and unread in tons of neglected Blue-Books. Then again the money has been carefully and laboriously collected, for the Committee were precluded by the terms of their reference from drawing on the purse of the taxpayer. This also made for economy.

Some criticisms have been heard at the delay which has occurred in the production of this volume. But it should be remembered that when the Inquiry started very little was accurately known about the Grouse either in health or in disease. As a member of the Scientific Staff said in a lecture before the Royal Institution: "In considering exceptions it is so immensely important to know the rule. In studying disease our starting-point should be the normal, the healthy; yet until lately no one has closely studied the healthy Grouse, and indeed it is almost impossible to find a normal Grouse, i.e., one free from parasites. A Grouse cannot express to us its feelings; the state of its tongue, the rate of its pulse, even its temperature tell us nothing because we have no norm and no means of estimating the extent to which a diseased Grouse has departed from the standards of a healthy bird. The nature of the numerous kinds of blood corpuscles, which alter in proportion so markedly in animals when they become parasitised, was but a few months ago quite unknown, the "blood count" uninvestigated; in fact the Inquiry started, as regards the cause and symptoms of the diseases which affect Grouse, practically behind scratch."

Further, the Committee were not in a position to retain the whole time of any one of their Scientific Staff with the single exception of the Field Observer. What work this staff have accomplished, and they have accomplished much, has been for the most part done in their spare time or during their brief holidays. Another factor that made for delay was that the Committee were not in a position to establish a central laboratory, and hence the actual investigations were carried on for a time in one place, and then after a break often of many weeks the threads were picked up in another. Much work was done at Cambridge, but at the London School of Tropical Medicine, at the Royal Scottish Museum, Edinburgh, at Frimley, at King's School in the Isle of Man, in the offices of the Field in London, in the gun-room at Beaufort, valuable investigations were also carried on. Further, from the necessity of examining absolutely fresh material, an improvised travelling laboratory had to be set up perhaps in a private sitting-room of a country hotel, perhaps in an outhouse of a Highland inn, but always under conditions which

vastly increased the difficulty of investigation, and made for delay.

Considering all these circumstances, the results now published do not seem unduly belated.

The Committee specially desire to record their thanks to the following gentle- men who have formed the Scientific Staff of the Inquiry, and to whose labours the results are due:—

Edward A. Wilson, M.B., F.Z.B., M.B.O.U., was appointed, in November 1905, principal Field Observer, Anatomist and Physiologist to the Inquiry, and devoted his whole time to the work till the autumn of 1910, when he joined Captain Scott's Antarctic Expedition as Scientific Director on the Terra Nova. It is difficult to speak highly enough of Dr. Wilson's services, for not only was he an indefatigable worker in the field, but his ornithological knowledge, his scientific training, and his artistic skill, have been of the utmost value in every branch of the Inquiry. Practically every Grouse which was submitted to the Committee for examination was dissected and reported on by Dr Wilson, and the results of these dissections, as shown in Appendix D, not only form a record of long and patient labour, but also provide an enormous mass of carefully arranged material which has been of great use to the Committee. Dr Wilson has written or aided in writing ten out of the first fourteen (Chapters of the Book, and has not only fully illustrated his own contributions, but has placed his artistic skill at the disposal of nearly all the other writers. In addition to his services as Field Observer and Physiologist, Dr Wilson conducted a series of experiments on live Grouse at the Committee's Observation Area whereby the results obtained by Dr Leiper, Dr Shipley and others were put to the test ; these experiments entailed some years of hard and patient work, and required the closest co-operation with the other members of the Scientific Staff. Dr Wilson's personal qualities secured for him the willing assistance alike of Local Correspondents and Scientific Staff, and went far to ensure whatever success the Committee has achieved.

A. E. Shipley, M.A., Hon. D.Sc, F.R.S., Master of Christ's College, Cambridge, and Reader in Zoology in the University of Cambridge, undertook in June 1905 to assist the Committee in the Scientific Departments of their research, especially in connection with the investigations of the ectoparasites and endoparasites of Grouse. Dr Shipley's services to the scientific side of the Inquiry have been as important as Dr Wilson's services to the natural history side. Dr Shipley has published the results of his labours in the Proceedings of the Zoological Society of London for 1909 in the following series of articles: (1) The Tapeworms (cestoda) of the Red Grouse; (2) The Threadworms (nematoda) of the Red Grouse; (3) The ectoparasites of the Red Grouse; (4) The Internal Parasites of birds allied to the Grouse. The first three of these papers are, by the courtesy of the Zoological Society of London, reprinted with minor alterations in the present Report. Dr Shipley has also acted as one of the Publishing Sub-Committee of the Inquiry, and has given much assistance in the revisal of the proofs and the preparation of Interim and Final Reports for the press.

R. F. Leiper, D.Sc, M.B., F.Z.S., Helminthologist to the London School of Tropical Medicine, was appointed in 1908 to help in the elucidation of certain difficult questions relating to the life history of the nematode worm Tricliostrongylus pergracilis, which in the opinion of the Committee is the main cause of mortality in adult Grouse. Dr Leiper devoted much time to the study of these questions, and to him is due the credit of having solved many of the problems connected with the development and bionomics of this important parasite. The result of his investigations are given in the present Report.

W. Bygrave and Percy H. Grimshaw assisted Dr Shipley by a prolonged and systematic search for the intermediate host of the Grouse tapeworms, and though the results were negative, the conscientious manner in which the search was conducted has enabled the Committee to claim that the question has been investigated as fully as was possible in the time at their disposal.

H. B. Fantham, D.Sc. Lond, B.A. Cantab., A.R.C.S., F.Z.S., Christ's College, Cambridge, Parasitologist to the Liverpool School of Tropical Medicine, formerly Assistant to the Quick Professor of Biology in the University of Cambridge, was appointed Protozoologist to the Inquiry in 1907, and since that date has made a careful study of the protozoal parasites which are found in the blood and alimentary tract of the Grouse. His researches have resulted in a most interesting series of discoveries, of which by far the most important from the Committee's point of view is that the Eimeria (Coccidium) avium frequently found in the alimentary tract of the Grouse is a frequent cause of death of young birds. Dr Fantham has followed up and fully described the life history of this parasite, whose presence in the intestine of the young Grouse was first pointed out by Dr. Leiper, and has published the results of his researches in the Proceedings of the Zoological Society of London for October 1910 in the following series of articles: (1) The Morphology and Life Hiatovy of Eimeria (Coccidium) avium: a Sporozoön causing a fatal disease among young Grouse; (2) Observations on the Parasitic Protozoa of the Red Grouse (Lagopus scoticus); (3) Experimental studies on Avian Coccidiosis, especially in relation to young Grouse, Fowls and Pigeons; (4) Observations on the Blood of Grouse. By the courtesy of the Zoological Society of London these articles are reprinted in the present Report.

C. G. Seligmann, M.B., then Pathologist to the Zoological Society of London, was appointed in 1906 to investigate the bacteriology of "Grouse Disease." He worked for the Committee till the end of 1907, when he left for Ceylon on a scientific expedition. The Committee is indebted to him for the discovery that the bacterial characters observed by Professor Klein as symptomatic of "Grouse Disease" were not in fact the pathological accompaniment of the mortality in Grouse as observed by the Committee. After Dr Seligmann went abroad his observations on this point were continued and confirmed by Dr Cobbett and Dr Graham Smith.

L. Cobbett, M.D., F.R.C.S., University Lecturer in Pathology, Cambridge, and G. S. Graham-Smith, M.D., University Lecturer in Hygiene, Cambridge, consented in 1909 to continue the work where Dr Seligmann had left off. They made an exhaustive investigation of the general pathology of "Grouse Disease" in all its forms, and the relation of the Bacillus coli of Professor Klein's "Grouse Disease" to the various pathological lesions which had come under the observation of the Committee. The results of their investigations were published in the Journal of Hygiene in June 1910, and, by the courtesy of Professor Nuttall, the Editor of that Journal, are reprinted in the present Report.

L. W. Sambon, M.D., gave considerable assistance to Dr Seligmann during the spring of 1907, and discovered a new leucocytozoon in the blood (L. Lovati).

H. Hammond Smith, M.B., Pathologist to the Field newspaper, has assisted the Committee both in the field and in the laboratory since the Inquiry was

commenced. He established and organised the Observation Area at Frimley in Surrey, and gave great assistance to the Committee in connection with the conduct of experiments at this Observation Area. He also assisted in the study of the question of the grits found in the gizzards of the Grouse and other game birds, and gave great help to the Committee in connection with the conduct of experiments at the Observation Area.

R. H. Rastall, M.A., F.G.S., Fellow and Lecturer of Christ's College, Cambridge, drew up an interesting report on the mineral constituents of gizzard grits in Grouse, and gave assistance in writing the article dealing with grits which appears in this Report. He also aided the work of publication by reading and correcting almost the whole of the proofs of this Report.

Percy H. Grimshaw, F. R.S.E., F.E.S., Assistant Keeper of the Natural History Department, Royal Scottish Museum, was appointed in 1909 to undertake the whole investigation of the insect life on the moors. He carried on and elaborated the work begun by Mr Fryer and Mr Hill, and not only prepared a complete list of the insects found on the moors, but also reported upon those eaten by the Grouse as shown by an examination of their crops and gizzards. The result of his work is published in the "Annals of Scottish Natural History" for July 1910, and in chapter iv. and Appendix E of the present Report. Mr Grimshaw also undertook the investigation of the habits and life history of the heather beetle (Lochmæa suturalis), and his article on this subject is included in the Report.

George C. Muirhead, B.Sc, acted as Field Observer from May to December 1905, and assisted in drawing up the pamphlet "Notes on the Grouse."

J. C. Fryer, B.A., Gonville and Caius College, Cambridge, was appointed in 1907 to make a report on the Insect Life of Grouse Moors. This Report has already appeared in the Interim Report of the Committee.

Alfred Hill was employed in 1908 to carry on the investigations already commenced by Mr Fryer.

A. S. Leslie, B.A., W.S. As soon as the Committee was officially appointed in 1905, one of their first acts was to nominate Mr Leslie as Secretary. During the six years that the Committee have sat he has continued to act in that capacity, and his duties have been both varied and arduous.

To him was entrusted the task of collecting the subscriptions, which formed the sole source of income for the Inquiry, and without which nothing could be done; the control of this Fund further rested in him. He also got together and organised the three hundred and sixty local correspondents, he drew up all the various tables, forms, etc., with which these correspondents were supplied, received the answers to the questions asked, collated and tabulated not only these answers but the verbal replies given at the several examinations of gamekeepers and other experts, which from time to time the Committee held. His correspondence amounted to many thousands of letters. Further, he assisted the Field Observer in many ways, especially in the preparation of statistics and the arrangement of tabular matter.

Mr Leslie wrote the "Notes on the Grouse," and has been in the main responsible for the preparation and seeing through the press both the Interim and the present (Final) Report ; the compiling of the appendices and the index, and the revision of the proofs, were largely his work.

To his knowledge of Scotland and of sport, and his professional training, the Committee owe many valuable suggestions as to the course the investigations have from time to time taken. They feel they cannot speak too highly of the self-sacrificing way he has thrown himself into the work, of his untiring energy, of his powers of organisation or of his adaptability and tact, which has done much to make the labours of not only the Committee but of all in any way associated with the Inquiry not only profitable but pleasurable.

The salary that the Committee have been able to otter to Mr Leslie can only be described as derisory. He has, in fact, received but the scantiest payment for the work he has done, and no compensation of any kind for the time he has taken from his profession and given to the Inquiry. But not only has he, like others, given time, skill and knowledge to further the cause of the investigation, but by his skilful husbandry of the limited resources available he has enabled the Committee to cover a wider area of research, and to prolong the time during which research was carried on to an extent whicli at first seemed impossible.

R. B. Fraser was appointed Assistant Secretary in October 1907, when it was found that the work of organisation and correspondence could not be conducted single handed by the Secretary. Mr Fraser has given valuable assistance with the general secretarial work, and also with the additional work entailed in connection with the preparation of the Report for the press.

In addition to those already mentioned the following have given the Committee much assistance in the revisal of proofs and in other ways: W. Berry, B.A., LL. B., M.B.O.U., who has been chiefly responsible for the Index; W. R. Ogilvie Grant, M.B.O.U., of the British Museum of Natural History; W. Eagle Clark, F.L.S., F.R.S.E., etc., Keeper of the Natural History Department of the Royal Scottish Museum; L. R. Sutherland, M.B., Professor of Pathology in the University of St Andrews ; Mrs E. A. Wilson, and the Hon. Gladys Graham Murray, F.Z.S.

An Abstract of Accounts is annexed, from which it may be seen how the income has been expended.[2]

The whole funds have now been exhausted in the work of investigation, and there is no balance available to meet the cost of publishing the results. This is to be regretted, as it will make it impossible to provide the supporters of the Inquiry with copies of the Report free of charge.

The thanks of the Committee are due to those moor-owners, shooting tenants, gamekeepers, and others who have gratuitously given their services as correspondents.

The Committee have to acknowledge with thanks the support it has received from its subscribers. A list of subscribers and the amount of their subscriptions is given in Appendix B.[3]

The Committee have also to acknowledge their indebtedness to the Zoological Society of London, which at the request of the Committee published in the Proceedings of the Society the articles on Ectoparasites and Endoparasites of Grouse by Dr A. E. Shipley ; the articles on the Protozoa and Blood of Grouse, by Dr H. B. Fantham ; and the article on the Plumage of Grouse, by Dr E. A. Wilson, comprising an important part of the scientific matter contained in this volume, which is reproduced here by consent of the Society. They have also to acknowledge their indebtedness to the Society for revising and editing the manuscript of Dr Wilson's contributions on the Plumage of the Grouse, in the absence of the author on the Antarctic Expedition.

The Committee also desire to acknowledge its indebtedness to the heads of the various Scientific Laboratories at Cambridge, where much of the research work was carried on; to the London School of Tropical Medicine who permitted Dr Lei per to assist in the investigation; and to the Directors and Staff of the Royal Scottish Museum, who assisted the Committee in Various ways during the whole period of the Inquiry.

August 1911.

  1. In the following chapters this worm is usually called Trichostrongylus pergracilis, but some writers have preserved Strongylus pergracilis, it is also at times called the Strongyle or the Strongyle worm. A synonym and a list of allied species are given by Dr Shipley on pp. 207 et seq.
  2. Vide vol. ii., Appendix C.
  3. Vide vol. ii., Appendix B.